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Ratcliffe, J. M., Fenton, M. B., & Shettleworth, S. J. (2006). Behavioral flexibility positively correlated with relative brain volume in predatory bats. Brain Behav Evol, 67(3), 165–176.
Abstract: We investigated the potential relationships between foraging strategies and relative brain and brain region volumes in predatory (animal-eating) echolocating bats. The species we considered represent the ancestral state for the order and approximately 70% of living bat species. The two dominant foraging strategies used by echolocating predatory bats are substrate-gleaning (taking prey from surfaces) and aerial hawking (taking airborne prey). We used species-specific behavioral, morphological, and ecological data to classify each of 59 predatory species as one of the following: (1) ground gleaning, (2) behaviorally flexible (i.e., known to both glean and hawk prey), (3) clutter tolerant aerial hawking, or (4) open-space aerial hawking. In analyses using both species level data and phylogenetically independent contrasts, relative brain size was larger in behaviorally flexible species. Further, relative neocortex volume was significantly reduced in bats that aerially hawk prey primarily in open spaces. Conversely, our foraging behavior index did not account for variability in hippocampus and inferior colliculus volume and we discuss these results in the context of past research.
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Gibson, B. M., Juricevic, I., Shettleworth, S. J., Pratt, J., & Klein, R. M. (2005). Looking for inhibition of return in pigeons. Learn Behav, 33(3), 296–308.
Abstract: We conducted four experiments in order to investigate whether pigeons' responses to a recently attended (i.e., recently pecked) location are inhibited. In Experiments 1 and 2, stimulus displays were similar to those used in studies of inhibition of return (IOR) with humans; responses to cued targets tended to be facilitated rather than inhibited. In Experiments 3 and 4, birds were presented with stimulus displays that mimicked clusters of small grains and were relatively localized, which should have been more appropriate for detecting IOR in pigeons. The results from these experiments again provided evidence for facilitation of responding to cued targets, rather than for IOR.
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Gibson, B. M., & Shettleworth, S. J. (2005). Place versus response learning revisited: tests of blocking on the radial maze. Behav Neurosci, 119(2), 567–586.
Abstract: Neurobiological and behavioral research indicates that place learning and response learning occur simultaneously, in parallel. Such findings seem to conflict with theories of associative learning in which different cues compete for learning. The authors conducted place+response training on a radial maze and then tested place learning and response learning separately by reconfiguring the maze in various ways. Consistent with the effects of manipulating place and response systems in the brain (M. G. Packard & J. L. McGaugh, 1996), well-trained rats showed strong place learning and strong response learning. Three experiments using associative blocking paradigms indicated that prior response learning interferes with place learning. Blocking and related tests can be used to better understand how memory systems interact during learning.
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Skov-Rackette, S. I., & Shettleworth, S. J. (2005). What do rats learn about the geometry of object arrays? Tests with exploratory behavior. J Exp Psychol Anim Behav Process, 31(2), 142–154.
Abstract: Six experiments using habituation of exploratory behavior tested whether disoriented rats foraging in a large arena encode the shapes of arrays of objects. Rats did not respond to changes in position of a single object, but they responded to a change in object color and to a change in position of 1 object in a square array, as in previous research (e.g., C. Thinus-Blanc et al., 1987). Rats also responded to an expansion of a square array, suggesting that they encoded sets of interobject distances rather than overall shape. In Experiments 4-6, rats did not respond to changes in sense of a triangular array that maintained interobject distances and angles. Shapes of object arrays are encoded differently from shapes of enclosures.
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Shettleworth, S. J., & Sutton, J. E. (2005). Multiple systems for spatial learning: dead reckoning and beacon homing in rats. J Exp Psychol Anim Behav Process, 31(2), 125–141.
Abstract: Rats homed with food in a large lighted arena. Without visual cues, they used dead reckoning. When a beacon indicated the home, rats could also use the beacon. Homing did not differ in 2 groups of rats, 1 provided with the beacon and 1 without it; tests without the beacon gave no evidence that beacon learning overshadowed dead reckoning (Experiment 1). When the beacon was at the home for 1 group and in random locations for another, there was again no evidence of cue competition (Experiment 2). Dead reckoning experience did not block acquisition of beacon homing (Experiment 3). Beacon learning and dead reckoning do not compete for predictive value but acquire information in parallel and are used hierarchically.
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Shettleworth, S. J. (2003). Memory and hippocampal specialization in food-storing birds: challenges for research on comparative cognition. Brain Behav Evol, 62(2), 108–116.
Abstract: The three-way association among food-storing behavior, spatial memory, and hippocampal enlargement in some species of birds is widely cited as an example of a new 'cognitive ecology' or 'neuroecology.' Whether this relationship is as strong as it first appears and whether it might be evidence for an adaptive specialization of memory and hippocampus in food-storers have recently been the subject of some controversy [Bolhuis and Macphail, 2001; Macphail and Bolhuis, 2001]. These critiques are based on misconceptions about the nature of adaptive specializations in cognition, misconceptions about the uniformity of results to be expected from applying the comparative method to data from a wide range of species, and a narrow view of what kinds of cognitive adaptations are theoretically interesting. New analyses of why food-storers (black-capped chickadees, Poecile Atricapilla) respond preferentially to spatial over color cues when both are relevant in a memory task show that this reflects a relative superiority of spatial memory as compared to memory for color rather than exceptional spatial attention or spatial discrimination ability. New studies of chickadees from more or less harsh winter climates also support the adaptive specialization hypothesis and suggest that within-species comparisons may be especially valuable for unraveling details of the relationships among ecology, memory, and brain in food-storing species.
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Gibson, B. M., & Shettleworth, S. J. (2003). Competition among spatial cues in a naturalistic food-carrying task. Learn Behav, 31(2), 143–159.
Abstract: Rats collected nuts from a container in a large arena in four experiments testing how learning about a beacon or cue at a goal interacts with learning about other spatial cues (place learning). Place learning was quick, with little evidence of competition from the beacon (Experiments 1 and 2). Rats trained to approach a beacon regardless of its location were subsequently impaired when the well-learned beacon was removed and other spatial cues identified the location of the goal (Experiment 3). The competition between beacon and place cues reflected learned irrelevance for place cues (Experiment 4). The findings differ from those of some studies of associative interactions between cue and place learning in other paradigms.
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Shettleworth, S. J., & Westwood, R. P. (2002). Divided attention, memory, and spatial discrimination in food-storing and nonstoring birds, black-capped chickadees (Poecile atricapilla) and dark-eyed juncos (Junco hyemalis). J Exp Psychol Anim Behav Process, 28(3), 227–241.
Abstract: Food-storing birds, black-capped chickadees (Poecile atricapilla), and nonstoring birds, dark-eyed juncos (Junco hyemalis), matched color or location on a touch screen. Both species showed a divided attention effect for color but not for location (Experiment 1). Chickadees performed better on location than on color with retention intervals up to 40 s, but juncos did not (Experiment 2). Increasing sample-distractor distance improved performance similarly in both species. Multidimensional scaling revealed that both use a Euclidean metric of spatial similarity (Experiment 3). When choosing between the location and color of a remembered item, food storers choose location more than do nonstorers. These results explain this effect by differences in memory for location relative to color, not division of attention or spatial discrimination ability.
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Hampton, R. R., & Shettleworth, S. J. (1996). Hippocampus and memory in a food-storing and in a nonstoring bird species. Behav Neurosci, 110(5), 946–964.
Abstract: Food-storing birds maintain in memory a large and constantly changing catalog of the locations of stored food. The hippocampus of food-storing black-capped chickadees (Parus atricapillus) is proportionally larger than that of nonstoring dark-eyed juncos (Junco hyemalis). Chickadees perform better than do juncos in an operant test of spatial non-matching-to-sample (SNMTS), and chickadees are more resistant to interference in this paradigm. Hippocampal lesions attenuate performance in SNMTS and increase interference. In tests of continuous spatial alternation (CSA), juncos perform better than chickadees. CSA performance also declines following hippocampal lesions. By itself, sensitivity of a given task to hippocampal damage does not predict the direction of memory differences between storing and nonstoring species.
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Hampton, R. R., & Shettleworth, S. J. (1996). Hippocampal lesions impair memory for location but not color in passerine birds. Behav Neurosci, 110(4), 831–835.
Abstract: The effects of hippocampal complex lesions on memory for location and color were assessed in black-capped chickadees (Parus atricapillus) and dark-eyed juncos (Junco hyemalis) in operant tests of matching to sample. Before surgery, most birds were more accurate on tests of memory for location than on tests of memory for color. Damage to the hippocampal complex caused a decline in memory for location, whereas memory for color was not affected in the same birds. This dissociation indicates that the avian hippocampus plays an important role in spatial cognition and suggests that this brain structure may play no role in working memory generally.
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